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The Spread of the Maritime Halophyte Cochlearia Danica Along Salted Continental Roads

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The Spread of the Maritime Halophyte Cochlearia Danica Along Salted Continental Roads Preslia 90: 23–37, 2018 23 A hitchhiker from the beach: the spread of the maritime halophyte Cochlearia danica along salted continental roads Šíření přímořského halofytního druhu Cochlearia danica podél solených silnic Réka F e k e t e1, Attila M e s t e r h á z y2, Orsolya V a l k ó3 & Attila M o l n á r V.1 1Department of Botany, University of Debrecen, Egyetem tér 1, H-4032 Debrecen, Hungary, email: [email protected], [email protected]; 2Directorate of Hortobágy National Park, H-4024, Debrecen Sumen u. 2, Hungary, email: [email protected]; 3MTA-DE Biodiversity and Ecosystem Services Research Group, H-4032 Debrecen Egyetem tér 1, Hungary, email: [email protected] Fekete R., Mesterházy A., Valkó O. & Molnár V. A. (2018): A hitchhiker from the beach: the spread of the maritime halophyte Cochlearia danica along salted continental roads. – Preslia 90: 23–37. The increase in road networks facilitates the dispersal of many species of plants along roadsides. In these special habitats, the use of deicing salt can provide suitable habitat conditions for the establishment of stress-tolerant halophytes. This study investigates the spread of an alien halophyte Cochlearia danica in continental Europe. This species is native to the Atlantic shores of Europe, has already spread in many countries along roadsides and was recently discovered in Hungary. We performed a literature review to track the European spread of this species, and investigated the Hungarian occurrences in detail. Then we determined the ability of this species to adapt to local soil conditions by means of soil analyses and germination tests using 19 different NaCl concentrations and alkaline soils. To estimate the rate of spread, we estimated the size of the four Hungarian populations in 2016 and 2017, and at the same time we measured the number of flowering stems, number of flowers in an inflorescence, number of seeds per fruit and seed mass. Cochlearia danica is recorded growing along roadsides in eight countries in continental Europe. Literature data indicate a rapid spread of this species along European roads, of 62–65 km/year. In Hungary this species is recorded at four roadside localities characterized by a high soil salt content. The relationship between NaCl concentration and percentage germination followed a sigmoidal curve. Germination tests revealed a significant negative effect of NaCl concentration on germination above 0.5% NaCl, but germination occurred even on extremely saline substrates with a 2% NaCl concentration. The area of the largest Hungarian population decreased by more than 99% and that of the second largest population increased by more than 30% between 2016 and 2017. Even though this species can adapt to high salinity in the soil, these rapid and marked changes in population size indicate that the population dynamics of this species may depend on interactions between the amount of local precipitation and soil type. Our study indicates that we should expect further occurrences of C. danica along roads. Keywords:Brassicaceae, deicing salt, NaCl, plant invasion, roadside verges, seed dispersal Introduction The extent of road networks has increased remarkably during the past century world- wide. The total length of global road networks is more than 64 million km and a further marked increase is projected for the near future (van der Ree et al. 2015). The length of doi: 10.23855/preslia.2018.023 24 Preslia 90: 23–37, 2018 paved roads in the European Union was recorded as 5,066,700 km in 2008 (Nicodeme et al. 2011). The mean size of areas without roads is 48 km2, which means that Europe has one of the densest road networks in the world (Ibisch et al. 2016). The rapidly growing density of roads has a negative effect on the occurrence of many taxa and decreases biodiversity (Findlay et al. 2000) and the typical negative effects include fragmentation and edge effects on surrounding habitats, thus the communities affected are becoming more vulnerable to negative effects, such as the spread of alien species, which are more able to become established in new habitats (Forman & Alexander 1998, Spellerberg 1998, Zwaenepoel et al. 2006). Roads, as ecological corridors, play a significant role in the spread of plant species (Tikka et al. 2001), long-distance dispersal of native and alien species by vehicles is a frequent phenomenon, but is more common among alien species (von der Lippe & Kowarik 2007). Propagule export from cities to rural and suburban environments is likely to be more frequent than from rural to urban areas, thus it could increase the incidence of plant invasions, as more propagules, with a higher representa- tion of seeds of alien species, were found along roads leading from urban areas (von der Lippe & Kowarik 2008). The spread of invasive species along roads is a worldwide phe- nomenon (Forman 2000, Gelbard & Belnap 2003, Kalwij et al. 2008, Essl et al. 2009, Follak et al. 2013, Skálová et al. 2017). Thus, roadsides are unique places, where often first occurrences of alien plants can be found. The construction and management of roads are usually associated with several types of anthropogenic disturbance, such as frequent mowing, deicing, use of herbicides, trampling by humans and vehicles, a range of pollut- ants and modified soils used for construction, the last of which may contain propagules of alien species (Šerá 2008, van der Ree et al. 2015). These all contribute to the spread of aliens, since native species are usually less able to adapt to the altered conditions in roadsides (Greenberg et al. 1997, Forman & Alexander 1998). In the case of lightweight seeds, dispersal may be facilitated by the air turbulence caused by cars (von der Lippe et al. 2013), and they may also travel long distances in mud attached to wheels (Ross 1986, Zwaenepoel et al. 2006), which can contain substantial numbers of seeds (Clifford 1959). For example, Schmidt (1989) sampled mud from a single car driven 15,000 km and found almost 4000 germinable seeds of more than 100 species. According to Ansong & Pickering (2013) more than 60,000 seeds were found in one ton of dry sludge collected from Australian cars. Machines used for mowingvergeshavealsobeenshowntotrans- port seeds (Strykstra et al. 1997, Vitalos & Karrer 2009). Use of deicing salt has become widespread in Europe during the last five decades. Although winter road maintenance practices differ between countries according to clima- tic conditions, in most European countries the most frequently used material for deicing is salt (mostly NaCl, with a small proportion CaCl2, rarely MgCl2; Houska 2007). The two countries using the largest quantities of salt are the United Kingdom and Germany (2200 and 2000 metric tons respectively; Houska 2007). Deicing salts have complex effects on the roadside environment; they infiltrate ground-water (Amrhein et al. 1992) and damage the leaves of many species of trees (Hofstra & Hall 1971). The increased soil salt content can cause osmotic stress for plants, alter soil pH and the availability of nutri- ents, thereby altering the species composition of roadside vegetation and facilitating the spread of halophytic species of plants (Davison 1971). Halophytes constitute approxi- mately 1% of the world’s flora and are able to survive and reproduce in environments where salt concentrations are at least 200 mM NaCl (Flowers & Colmer 2008). Obligate Fekete et al: Spread of Cochlearia danica along roads 25 halophytes thrive best in saline circumstances (Flowers & Colmer 2008). Facultative halophytes are poor competitors and restricted to extreme environmental conditions (Barbour 1978). Spread of a maritime halophyte species, Plantago coronopus, was first detected in Hungary in 2013 (Schmidt et al. 2014), and further occurrences were recorded in 2014, 2015 (Kovács & Lengyel 2015) and 2016 (Schmidt et al. 2016), which indicate an east- ward spread of this species. Spread of native halophytes, such as Plantago maritima (Barina 2007), Puccinellia distans (Király & Hohla 2015), Limonium gmelinii, Atriplex tatarica, Podospermum canum (Schmotzer 2015), Spergularia media, S. marina and Bupleurum tenuissimum (Schmidt et al. 2016) has also been documented. The motivation for this study was the recent discovery of an Atlantic coastal halophyte plant species, Cochlearia danica (Danish scurvygrass) in Hungary in 2016 (Molnár V. & Löki 2016). This discovery offered a unique opportunity to track the first stages of a potential plant invasion. Here, we analyse the spread of this species in central Europe, and assess its ability to bypass geographical, environmental and reproductive barriers during its spread. We focused on the following stages of a potential invasion: (i) Transport. We tracked the dispersal of this species through continental Europe based on reports in the literature and investigated in detail its recent occurrences in Hungary based on a comprehensive field survey. (ii) Colonization. We examined soil parameters associated with the Hungar- ian roadside habitats of C. danica and tested the effect of soil salt content on its germinability in an in vitro germination experiment. (iii) Establishment. We estimated its ability to spread by studying its population dynamics and reproductive traits. Finally we estimated its invasion potential in continental Europe. Material and methods Surveying the spread of Cochlearia danica through Europe To estimate the speed of spread of this species in continental Europe, we searched the lit- erature using Google Scholar and the following keywords: the name of the species ‘Cochlearia danica’ AND ‘roadside’ OR ‘road verge’. This returned 94 hits. After scan- ning these 94 papers, the number of studies dealing with roadside occurrences of Cochlearia danica was 13. Among these 13 papers, we found only one study dealing with continental Europe, and three more in the reference lists of the other 12 papers. We did not include the data on the occurrences of C. danica in the British Isles because this species there is already widespread along inland roads.
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